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PubMed2003

Isoflavonoid biosynthesis and accumulation in developing soybean seeds.

Isoflavonoids are biologically active natural products that accumulate in soybean seeds during development. The amount of isoflavonoids present in soybean seed is variable, depending on genetic and environmental factors that are not fully understood. Experiments were conducted to determine whether isoflavonoids are synthesized within seed tissues during development, or made in other plant organs and transported to the seeds where they accumulate. An analysis of isoflavonoids by HPLC detected the compounds in all organs of soybean plant, but the amount of isoflavonoids present varied depending on the tissue and developmental stage. The greatest concentrations were found in mature seeds and leaves. The 2-hydroxyisoflavanone synthase genes IFS1 and IFS2 were studied to determine their pattern of expression in different tissues and developmental stages. The highest level of expression of IFS1 was observed in the root and seed coat, while IFS2 was most highly expressed in embryos and pods, and in elicitor-treated or pathogen-challenged tissues. Incorporation of radiolabel into isoflavonoids was observed when developing embryos and other plant organs were fed with [(14)C]phenylalanine. Embryos excised from developing soybean seeds also accumulated isoflavonoids from a synthetic medium. A maternal effect on seed isoflavonoid content was noted in reciprocal crosses between soybean cultivars that differ in seed isoflavonoids. From these results, we propose that developing soybean embryos have an ability to synthesize isoflavonoids de novo, but that transport from maternal tissues may in part contribute to the accumulation of these natural products in the seed.

Carbon Radioisotopes↗
PubMed1990

Pattern of Variations in Abscisic Acid Content in Suspensors, Embryos, and Integuments of Developing Phaseolus coccineus Seeds.

Free abscisic acid (ABA) content in suspensors, embryos, and integuments was determined during seed development of Phaseolus coccineus. A highly specific and sensitive solid-phase radioimmunoassay based on a monocional antibody raised against free (S)-ABA was used for ABA quantification. Very small amounts of ABA were detected in the suspensor during initial stages of development; later two peaks of ABA occurred. Levels of ABA in the embryo and integument show a coincident triphasic distribution: two maxima in ABA content occurred when the embryo was 11 to 12 and 15 to 16 millimeters in length; later, when the embryo was 19 to 20 millimeters long, a further increase was observed. The role of ABA in runner bean seeds is discussed in relation to the development of the different seed tissues.

Journal Article↗
PubMed2006

Pinoresinol-lariciresinol reductase gene expression and secoisolariciresinol diglucoside accumulation in developing flax (Linum usitatissimum) seeds.

The transcription activity of the pinoresinol-lariciresinol reductase (PLR) gene of Linum usitatissimum (so-called LuPLR), a key gene in lignan synthesis, was studied by RT-PCR and promoter-reporter transgenesis. The promoter was found to drive transcription of a GUSint reporter gene in the seed coats during the flax seed development. This fitted well with the tissue localization monitored by semi-quantitative RT-PCR of LuPLR expression. Accumulation of the main flax lignan secoisolariciresinol diglucoside was coherent with LuPLR expression during seed development. This three-way approach demonstrated that the LuPLR gene is expressed in the seed coat of flax seeds, and that the synthesis of PLR enzyme occurs where flax main lignan is found stored in mature seeds, confirming its involvement in SDG synthesis.

Base Sequence↗
PubMed1989

Carbon Partitioning among Leaves, Fruits, and Seeds during Development of Phaseolus vulgaris L.

Development of vegetative and floral buds was found to be a key factor in establishing the way carbon is distributed among growing leaves and fruits in Phaseolus vulgaris L. plants. Leaves emerged principally during a period 14 to 32 days after planting while flowers were produced during a 10- to 12-day period near the end of leaf emergence. Timing of anthesis established the sigmoidal time course for dry weight accumulated by the composite of all fruits on the plant. During the first 12 days following anthesis, fruit growth mainly consisted of elongation and dry weight accumulation by the pod wall. Thereafter, seed dry weight increased for about 1 week, decreased markedly for several days, and then increased again over the next 2 weeks. Accumulation of imported carbon in individual seeds, measured by steady-state labeling, confirmed the time course for dry weight accumulation observed during seed development. Seed respiration rate initially increased rapidly along with dry weight and then remained nearly steady until seed maturation. A number of developmental events described in the literature coincided with the different phases of diauxic growth. The results demonstrated the feasibility of relating current rates of carbon import in individual seeds measured with tracer (14)C to the rates of conversion of imported sucrose and use of the products for specific developmental processes. The resulting data are useful for evaluating the roles of conversion and utilization of imported sucrose in regulating import by developing seeds.

Journal Article↗
PubMed1995

Microsomal Lyso-Phosphatidic Acid Acyltransferase from a Brassica oleracea Cultivar Incorporates Erucic Acid into the sn-2 Position of Seed Triacylglycerols.

Developing seeds from Brassica oleracea (L.) var botrytis cv Sesam were examined for the ability to biosynthesize and incorporate erucic acid into triacylglycerols (TAGs). Seed embryos at mid-development contained a high concentration of erucic acid in diacylglycerols and TAGs, and substantial levels were also detected in free fatty acids, acyl-coenzyme A (CoA), phosphatidic acid, and phosphatidylcholine. Homogenates and microsomal fractions from seeds at mid-development produced [14C]eicosenoyl- and [14C]erucoyl-CoAs from [14C]oleoyl-CoA in the presence of malonyl-CoA and reducing equivalents in vitro. These fatty acids were incorporated into TAGs via the Kennedy pathway. However, unlike most Brassicaceae, the B. oleracea was able to insert significant erucic acid into the sn-2 position of TAGs. It was shown that the lyso-phosphatidic acid acyltransferase (LPAT) incorporated erucic acid into the sn-2 position of lyso-phosphatidic acid. The erucoyl-CoA:LPAT activity during seed development and the sn-2 erucic acid content of the TAG fraction in mature seed were compared to those in B. napus, Tropaeolum majus, and Limnanthes douglasii. There was a correlation between the in vitro erucoyl-CoA:LPAT activity and the sn-2 erucic acid content in seed TAGs. To our knowledge, this is the first member of the Brassicaceae reported to have an LPAT able to use erucoyl-CoA. This observation has important implications for efforts being made to increase the erucic acid content in B. napus, to supply strategic industrial feedstocks.

Journal Article↗
PubMed2006

Identification and functional expression of a type 2 acyl-CoA:diacylglycerol acyltransferase (DGAT2) in developing castor bean seeds which has high homology to the major triglyceride biosynthetic enzyme of fungi and animals.

Seed oil from castor bean (Ricinus communis) contains high amounts of hydroxy fatty acid rich triacylglycerols (TAGs) that can serve as raw material for production of bio-based products such as nylon, cosmetics, lubricants, foams, and surfactants. Diacylglycerol acyltransferase (DGAT) catalyses the terminal reaction in the acyl-CoA dependent Kennedy pathway of triglyceride biosynthesis. There is still some debate whether there are three or four enzymes in yeast that have DGAT activity and catalyse the synthesis of TAG but of these the DGAT2 homologue Dga1 contributes in a major way to TAG biosynthesis. Here we report on the cloning of a cDNA for DGAT2 from castor bean and prove its biological activity following expression in yeast and enzymatic assays using diricinolein as the acceptor and ricinoleoyl-CoA as the donor. Previous reports of DGAT in castor have focussed on DGAT1 which has little amino acid sequence homology to DGAT2. Expressional studies demonstrate that DGAT2 is 18-fold more highly expressed in seeds than in leaves and shows temporal specific expression during seed development. In contrast, DGAT1 shows little difference in expression in seeds versus leaves. We conclude that in castor bean DGAT2 is more likely to play a major role in seed TAG biosynthesis than DGAT1.

Amino Acid Sequence↗
PubMed1993

The gene encoding a calcium-dependent protein kinase located near the sbe1 gene encoding starch branching enzyme I is specifically expressed in developing rice seeds.

A gene (spk) encoding a Ca(2+)-dependent protein kinase (SPK) is located in the region immediately upstream of the sbe1 gene encoding a starch branching enzyme. The spk gene is specifically expressed in developing seeds and its expression pattern is very similar to those of genes encoding starch-synthesizing enzymes such as sbe1 and waxy, seed lipid-synthesizing enzymes, as well as genes encoding seed storage proteins. A full-length spk cDNA was isolated from a cDNA library constructed from developing seeds. The deduced amino acid sequence showed that SPK has a high degree of homology to soybean and carrot Ca(2+)-dependent protein kinase, both of which contain calmodulin domains. The calmodulin domain, as well as the catalytic subdomain consensus regions of protein kinases are highly conserved in SPK. These results suggest that a tissue- and stage-specific protein kinase, SPK, is involved in the synthesis of seed storage compounds during seed development. They also strongly suggest that Ca2+ is required for seed development.

1,4-alpha-Glucan Branching Enzyme↗
PubMed1985

Sucrose Concentration at the Apoplastic Interface between Seed Coat and Cotyledons of Developing Soybean Seeds.

The apoplastic sucrose concentration at the interface between cotyledons and surrounding seed coats of developing soybeans (Glycine max L. Merr. cv Wye) was found by three indirect methods to be in the range of 150 to 200 millimolar. This is an order of magnitude higher than has been reported elsewhere for soybean. It was also higher than the overall sucrose concentrations in the cotyledons and seed coats, each of which was approximately 90 millimolar. By defoliating plants 24 hours before measurement, both the overall sucrose concentration in the cotyledons and the interfacial apoplastic sucrose concentration were reduced by three-fourths. However, there was no day/night difference in overall tissue sucrose concentration of cotyledons or seed coats from intact plants suggesting the existence of a homeostatic mechanism compensating for the diurnal photosynthetic cycle. About 7 hours were required for a tritiated polyethylene glycol-900 solution to fully permeate developing cotyledons (from approximately 220 milligram fresh weight embryos), implying high diffusion resistance through the tissue.These results indicate that a high interfacial sucrose concentration may exist in vivo. They suggest that the saturable carrier-mediated component of sucrose uptake may be of little physiological significance in the outermost cell layers of the cotyledons.

Journal Article↗
PubMed2006

Regulation of carotenoid and ABA accumulation during the development and germination of Nicotiana plumbaginifolia seeds.

Abscisic acid (ABA) is derived from epoxycarotenoid cleavage and regulates seed development and maturation. A detailed carotenoid analysis was undertaken to study the contribution of epoxycarotenoid synthesis to the regulation of ABA accumulation in Nicotiana plumbaginifolia developing seeds. Maximal accumulation of xanthophylls occurred at mid-development in wild type seeds, when total ABA levels also peaked. In contrast, in ABA-deficient mutants xanthophyll synthesis was delayed, in agreement with the retardation in seed maturation. Seed dormancy was restored in mutants impaired in the conversion of zeaxanthin into violaxanthin by zeaxanthin epoxidase (ZEP), by the introduction of the Arabidopsis AtZEP gene under the control of promoters inducing expression during later stages of seed development compared to wild type NpZEP, and in dry and imbibed seeds. Alterations in the timing and level of ZEP expression did not highly affect the temporal regulation of ABA accumulation in transgenic seeds, despite notable perturbations in xanthophyll accumulation. Therefore, major regulatory control of ABA accumulation might occur downstream of epoxycarotenoid synthesis.

Abscisic Acid↗
PubMed1978

The biosynthesis of linoleate from oleoyl-CoA via oleoyl-phosphatidylcholine in microsomes of developing safflower seeds.

Microsomes prepared from developing safflower seeds rapidly desaturated added [14C]oleoyl-CoA in the presence of NADH. The distribution of [14C]oleate and [14C]linoleate between different lipid classes revealed that phosphatidylcholine was labelled with [14C]linoleate before any other lipid class investigated. Considerable desaturation continued after the disappearance of [14C]oleoyl-CoA in the reaction mixture. No [14C]linoleoyl-CoA could be detected. Incubation in the absence of added NADH caused a rapid disappearance of [14C]oleoyl-CoA by incorporation into triacylglycerols and phosphatidylcholine and also by release as [14C]oleic acid. Upon subsequent addition of NADH, [14C]linoleate was formed to the same extent as when NADH was present at the onset of the incubation. These data are contradictory to the previously assumed pathway for linoleic acid biosynthesis in developing safflower seeds, claiming oleoyl-CoA as the direct substrate and linoleoyl-CoA as the primary product. However, our data corroborate published results on Chlorella, Candida, Torulopsis and on developing pea leaves, which strongly suggest that oleoyl-CoA is first incorporated into phosphatidylcholine and then desaturated to linoleoyl-phosphatidylcholine.

Acyl Coenzyme A↗
PubMed2003

A novel C-terminal proteolytic processing of cytosolic pyruvate kinase, its phosphorylation and degradation by the proteasome in developing soybean seeds.

Cytosolic pyruvate kinase (ATP:pyruvate 2-O-phosphotransferase, EC 2.7.1.40) is an important glycolytic enzyme, but the post-translational regulation of this enzyme is poorly understood. Sequence analysis of the soybean seed enzyme suggested the potential for two phosphorylation sites: site-1 (FVRKGS220DLVN) and site-2 (VLTRGGS407TAKL). Sequence- and phosphorylation state-specific antipeptide antibodies established that cytosolic pyruvate kinase (PyrKinc) is phosphorylated at both sites in vivo. However, by SDS-PAGE, the phosphorylated polypeptides were found to be smaller (20-51 kDa) than the full length (55 kDa). Biochemical separations of seed proteins by size exclusion chromatography and sucrose-density gradient centrifugation revealed that the phosphorylated polypeptides were associated with 26S proteasomes. The 26S proteasome particle in developing seeds was determined to be of approximately 1900 kDa. In vitro, the 26S proteasome degraded associated PyrKinc polypeptides, and this was blocked by proteasome-specific inhibitors such as MG132 and NLVS. By immunoprecipitation, we found that some part of the phosphorylated PyrKinc was conjugated to ubiquitin and shifted to high molecular mass forms in vivo. Moreover, recombinant wild-type PyrKinc was ubiquitinated in vitro to a much greater extent than the S220A and S407A mutant proteins, suggesting a link between phosphorylation and ubiquitination. In addition, during seed development, a progressive accumulation of a C-terminally truncated polypeptide of approximately 51 kDa was observed that was in parallel with a loss of the full-length 55 kDa polypeptide. Interestingly, the C-terminal 51 kDa truncation showed not only pyruvate kinase activity but also activation by aspartate. Collectively, the results suggest that there are two pathways for PyrKinc modification at the post-translational level. One involves partial C-terminal truncation to generate a 51 kDa pyruvate kinase subunit which might have altered regulatory properties and the other involves phosphorylation and ubiquitin conjugation that targets the protein to the 26S proteasome for complete degradation.

Amino Acid Sequence↗
PubMed1981

Adenylate and nicotinamide nucleotides in developing soybean seeds during seed-fill.

Profiles of adenylate and nicotinamide nucleotides in soybean seeds were determined during seed-fill. The ATP content per seed increased during the early seed-filling stages to a level of 10 to 12 micrograms per seed. Seed ATP decreased after 40 days of development and reached its lowest level of less than 1 microgram at maturity. The ATP:ADP ratios were relatively constant at all seed development stages. Sharp increases in AMP levels during the late seed-fill stages were paralleled with a disappearance of ATP and ADP pools resulting in a reduced seed energy charge. Energy charge varied from the highest value of 0.78 at mid-seed-fill to less than 0.10 at maturity.Of the oxidized (NAD, NADP) and reduced (NADH, NADPH) nicotinamide nucleotide forms, NAD was the most abundant. Levels as high as 17.5 micrograms per seed were observed during the mid-seed-filling stages. NADP was found almost exclusively in the reduced form with a NADP: NADPH ratio of less than 0.35, whereas the reverse was noted for NAD which was found mainly in the oxidized form with a NAD:NADH ratio in the range of 5 to 25. NADP was detected in low concentrations compared to the other adenylate and nicotinamide nucleotides. The nicotinamide redox charge defined as (NADH + NADPH)/(NAD + NADH) + (NADP + NADPH) was calculated to express the state of the energy balance between the oxidized and reduced nicotinamide nucleotide forms. The nicotinamide redox charge varied between 0.15 and 0.30 during seed development and was significantly lower than that found for the adenylate energy charge.

Journal Article↗
PubMed2004

Pulsing Cl- channels in coat cells of developing bean seeds linked to hypo-osmotic turgor regulation.

Seed coat cells in the developing seeds of grain legumes release nutrients to the developing embryo. This occurs into an apoplastic space that separates the maternal (seed coat) and filial (embryo) generations. Protoplasts of seed coat cells from coats of Phaseolus vulgaris L. seeds were isolated and whole-cell current across their plasma membranes was characterized using the patch-clamp technique. A pulsing inward current that displayed a spontaneous activation and voltage-dependent inactivation was observed. The frequency and magnitude of the current pulses were positively dependent on cytoplasmic Cl(-) concentrations and independent of external cations. The pulse current was inhibited by DIDS and La(3+), but not by Gd(3+). Single channel events (conductance=18 pS) could be identified with the inactivating phase of the pulses. Together, these findings are consistent with the current being carried by a burst of Cl(-) efflux through Cl(-)-permeable channels that activate almost simultaneously. Neomycin caused a reversible inhibition of the pulsed current, suggesting that its activation is likely to be modulated by an IP(3)-dependent intracellular Ca(2+) release. The pharmacological profiles of Cl(-) efflux from excised seed coats were comparable with those of the Cl(-) channels in the whole cell configuration, suggesting that the Cl(-) channels may underpin Cl(-) efflux from the seed coats. Efflux of Cl(-) from the seed coats was also stimulated by hypo-osmotic treatment as was the frequency and magnitude of Cl(-) channel in whole-cell patch clamp experiments. This implies that the Cl(-) channels responsible for the pulsed Cl(-) currents are likely to be a component of the turgor-regulatory mechanism in developing bean seeds.

4,4'-Diisothiocyanostilbene-2,2'-Disulfonic Acid↗
PubMed2005

Development and composition of the seeds of nine genotypes of the Medicago truncatula species complex.

The seed development and composition of Medicago truncatula Gaertn., the new model plant for grain legumes, was studied using nine genotypes of the species complex: M. truncatula-Medicago littoralis (M. truncatula). The seed development of M. truncatula was very similar to that of other legumes, the only notable exception being the presence, in the mature seed, of an endosperm layer that is absent in grain legumes. During early embryogenesis and until mid-maturation, transient storage of starch occurred in the seed coat and embryo. This temporary storage probably contributed to the early development of the embryo and reserve synthesis. During maturation the synthesis and accumulation of proteins and oil took place at quasi-constant rates. Conversely oligosaccharides, mainly stachyose, were synthesised only during late maturation and at the beginning of desiccation. Proteins represented the major class of storage compounds and their average amino acid composition was found to be very close to that of pea and robust in various environmental conditions. Similar compositions between the two species and other grain legumes were also found for the fatty acids and the soluble sugars; most of these characters varied depending on the various environmental conditions used for seed production. All these similarities fully justify the use of M. truncatula as a model plant for genomic approaches to grain legume improvement. The major difference between M. truncatula seeds and European grain legume seeds resides in the nature of their carbon storage namely triacylglycerides for M. truncatula and starch for pea and faba bean.

Amino Acids↗
PubMed1992

Frost, Abscisic Acid, and Desiccation Hasten Embryo Development in Brassica napus.

Seed development in canola (Brassica napus) following a mild nonlethal freeze was examined with respect to abscisic acid (ABA) levels, desiccation, and expression of LEA.76 and isocitrate lyase (ICL) transcripts. Plants with seed of 70 and 55% moisture contents were frozen to -5 degrees C for 3 hours, and seed development followed after thawing. In addition, similar processes were compared during induction of extreme desiccation tolerance by application of ABA in Brassica microspore-derived haploid embryos in culture. A mild freeze/thaw caused a premature switch in seed developmental direction from predesiccation to desiccation as indicated by an immediate and accelerated loss of seed moisture to levels similar to the mature seed in 7 instead of 35 days, and by elevated ABA levels and induction of low levels of LEA.76 and ICL transcripts. Similarly, addition of ABA to haploid embryos in culture resulted in the induction of desiccation tolerance and low levels of late embryogenesis-abundant (LEA) but not ICL transcripts. In contrast, normal seed development and desiccation of ABA-treated (desiccation-tolerant) embryos resulted in the induction of ICL and very high levels of LEA.76 transcripts. Similarly, desiccation of control (desiccation-sensitive) embryos resulted in very high levels of LEA.76 transcripts. These results indicate that although LEA-type proteins have been implicated in the development of desiccation tolerance, high transcript levels of LEA.76 were not observed in the induction of desiccation tolerance either by a hastening of the maturation process in the developing Brassica seed, or by the exogenous application of ABA to Brassica haploid embryos in culture.

Journal Article↗
PubMed1969

Two additional phosphorylases in developing maize seeds.

Two additional phosphorylases (III and IV) have been detected in developing seeds of maize. Phosphorylase IV is found only in the embryo (with scutellum). It is also present in the embryo of the germinating seed where its activity is 90-fold greater than the activity in the developing embryo 22 days after pollination. Phosphorylase IV is eluted from a DEAE-cellulose column in the same fraction as phosphorylase I of the endosperm, and the 2 enzymes are similar in many respects. Phosphorylase IV is distinguished from phosphorylase I by electrophoretic mobility, by pH optimum, and because its properties are not affected by the shrunken-4 mutation. Phosphorylase III is found both in the endosperms and embryos of developing seeds. Activity for this enzyme is not detected in crude homogenates nor eluates from a DEAE-cellulose column apparently because it complexes with a non-dialyzable, heat-labile inhibitor. High activity is found after protamine sulfate fractionation. Phosphorylase III is bound to protamine sulfate and is then removed by washing with 0.3 m phosphate buffer. Phosphorylase III activity in the endosperm is not detectable 8 days after pollination but is present 12 days after pollination. Phosphorylase III differs from phosphorylases I, II, and IV in several respects-pH optimum, pH-independent ATP inhibition, time of appearance in the endosperm, and because purine and pyrimidine nucleotides are equally inhibitory. In common with phosphorylase II, phosphorylase III apparently does not require a primer to initiate the synthesis of an amylose-like polymer.

Glucosyltransferases↗
PubMed2002

Arabidopsis mutants deficient in diacylglycerol acyltransferase display increased sensitivity to abscisic acid, sugars, and osmotic stress during germination and seedling development.

Arabidopsis seeds store triacylglycerol (TAG) as the major carbon reserve, which is used to support postgerminative seedling growth. Diacylglycerol acyltransferase (DGAT) catalyzes the final step in TAG synthesis, and two isoforms of DGAT have previously been identified in Arabidopsis. It has been shown that DGAT1 plays an important role in seed development because Arabidopsis with mutations at the TAG1 locus accumulate less seed oil. There is also evidence showing that DGAT1 is active after seed germination. The aim of this study is to investigate the effect of mutations of DGAT1 on postembryonic development in Arabidopsis. We carried out detailed analyses of two tag1 mutants in different ecotypic backgrounds of Arabidopsis. Results show that during germination and seedling growth, seed storage TAG degradation was not affected in the tag1 mutants. However, sugar content of the mutant seedlings is altered, and activities of the hexokinases are significantly increased in the tag1 mutant seedlings. The tag1 mutants are also more sensitive to abscisic acid, glucose, and osmotic strength of the medium in germination and seedling growth.

Abscisic Acid↗